Taxonomic description and documentation of morphology of the invasive Mediterranean
Pararotalia calcariformata
Superfamily ROTALIOIDEA Ehrenberg, 1839
Family ROTALIIDAE Ehrenberg, 1839
Subfamily PARAROTALIINAE Reiss, 1963
Genus Pararotalia Le Calvez, 1949
Pararotalia calcariformata McCulloch, 1977
Not Pararotalia spinigera Le Calvez, 1949, emend. by Loeblich and Tappan (1957), Plate 4, Figs
1-3, [middle Eocene, Lutetian, Calcarie Grossier, France]
Not Pararotalia spinigera Le Calvez, 1949, Hottinger et al. (1991) Plate 1 Figs. 2, 5-8 [middle
Eocene, Lutetian, Grignon, Paris Basin, France]
Pararotalia (?) calcariformata McCulloch, 1977, McCulloch (1977), Plate 177, Figs 10-11,
[Recent, type locality Colombo Bay, shallow waters, Station 616, off West Nole Island,
Australia, shallow waters]
Pararotalia calcariformata McCulloch 1977, Loeblich and Tappan (1994), Plate 367, Figs. 10-13,
[Hypotype, Southern Timor Sea, Figs 11-13 sample V-347 at 58.52 m, Recent, North of
Bathurst Island, southeast Timor Sea, 1961 Sahul Shelf Cruise II Stranger]
Pararotalia spinigera Le Calvez 1949, Reinhardt et al. (1994), Plate 2, Figs 11-12. [Late
Holocene, Recent, CT/Grab 3, 9 m, Caesarea, Israel]
Eponides repandus Fichtel et Moll, 1798, Yanko et al. (1994), Plate 2, Figs. 1-9 [Recent, Haifa
Bay, Station 27, 12 meters]
Pararotalia spinigera Le Calvez, 1949, Arieli et al. (2011), Plate 6, Figs. 9-12 [Recent, Hadera,
Israel]
Pararotalia (?) calcariformata McCulloch, 1977, Meriç et al. (2013), Plate 1 Figs. 1-11, not 12
[Recent, Hatay, Turkey, 3-8 meters]
Original species description: Test free, calcareous, auriculate, biconvex, trochospiral,
periphery lobulated, craniate with usually one short spine per chamber, dorsally wall rather
smooth semi-hyaline, finely perforate; dorsal side evolute showing less than three whorls of
slightly inflated subrhomboidal chambers gradually increasing in size; sutures dorsally limbate
darker hyaline flush to depressed curved bands. Ventral side involute with prominent umbilical,
raised nodulous, hyaline to umbilical plug surrounded by nodulous umbilical flaps, usually seven
chambers in the last formed whorl; keel shallow hyaline acute with most of the tapering
peripheral chambers forming a single short angular hyaline spine extending out in posteriorly
from anterior half of each chamber of last formed whorl; sutures ventrally rather broad limbate
depressed radiate; aperture interiomarginal extraumbilical umbilical with a narrow depressed rim
outlining a low arch beginning at the close to the periphery, no tooth plate visible due to excess
shell on most specimens.
Comments: The Levantine Pararotalia examined by us (see Plate 1 and 2) reveals a significant
morphological variability, which is accentuated by ontogeny. Juvenile forms (Plate 2) typically
possess trapezoidal chambers and single pseudospines (occasionally two pseudospines per
chamber) extending from most chambers of the last whorl, occasionally positioned near the
septal face. Chambers of the last whorl in fully developed adults are usually petaloid and lack
the peripheral pseudospines. The peripheral margins and keel of the umbilical side in both
juveniles and adults are covered with numerous short pustules, found also in P. spinigera Le
Calvez 1949. The studied specimens from Israel reveal two pore structures that have not been
documented previously: micropores and larger circular to elliptical pore mounds on the surface
of both sides of the test in juvenile and adult forms. The chamber wall of Pararotalia sp. is highly
transparent, similar to other foraminifera known to host symbionts such as other Calcarinids. It
has been suggested that such a wall structure can be seen as an adaptation to photosymbiosis
(Leutenegger 1984). The range of variability in the examined populations include all
morphologies figured by Meriç et al. (2013). These authors note the lack of spines in their
specimens, but we believe they do so in contrast to species of Pararotalia with more prominent
spines, such as P. stellata. Not all of the Levantine specimens studied by us had spines (Plate 1,
1-3). Also, we note that a specimen pictured by Meriç et al. (2013) in their Figure 2, image 4a,
seems to possess spines of a similar shape and extent as in the studied population from Israel
(Plate 1, 4). Therefore, we conclude that the populations from Israel and Hatay, Turkey, are
morphologically overlapping. Following Meriç et al. (2013) we conclude that the Levantine
Pararotalia is morphologically distinct from the concept of P. spinigera. The latter species has a
lobate periphery without keel, which is a distinct and persistent feature of the Levantine
population (Plate 1, 2-4, 6). In addition, in the Levantine population the umbilical sutures are so
deeply incised that the walls of adjacent chambers become partly disconnected, forming deep
interlocular spaces (Plate 1, 1, 3, 4, 6). This character is indicated in the illustration of the type
material of P. calcariformata (McCulloch 1977). The particular character also clearly
distinguishes this species from the youngest fossil species of the genus in the Mediterranean,
described by Mancin et al. (2000) from Pliocene sediments in northern Italy as Pararotalia
padana. The Pliocene P. padana also show considerable variability, similar overall shape and
presence of spines. On the other hand, P. padana has more chambers in the last whorl, more
lobate periphery and completely lacks the interlocular spaces between successive chambers
that are observed on the umbilical side of many specimens in P. calcariformata. In addition, P.
padana does not show the distinct depressed area near the periphery on the umbilical side of
the shell (see specimen in Plate 1, Figure 3d) and appears to lack well developed nodes that
surround the chambers of P. calcariformata on the umbilical side.
Plate A
Late ontogenetic stages of Pararotalia calcariformata
1. Adult specimen with dextral coiling. a-b spiral views, note the slightly raised sutures, a
rounded peripheral outline, the spinose peripheral keel, and the central thickening of the early
whorls. b. Details showing densely perforated pore fields between imperforate septal sutures. cd Umbilical (ventral) views, note the interlocular space between consecutive chambers, the
presence of umbilical plug fused with the umbilical wall of the surrounding chambers, the thick
marginal nodes on early chambers, and the hirsute peripheral surface with numerous pustules.
2. a. Spiral view of fully developed adult specimen (sinistral coiling), with nine chambers in the
last whorl. b-c Lateral view of adult specimen showing interiomarginal aperture
3. Adult specimen with dextral coiling. a-b Spiral view with a frequent well-developed pore
mounds. c-d Umbilical view showing umbilical plug, large nodes along the edges of the sutures
and chamber surface with well-developed pore mounds. e. Lateral view showing interiomarginal
aperture.
4. Adult specimen with dextral coiling. a-b Spiral views showing nodose ornamentation,
peripheral spines on early chambers, and microperforate surface of the ultimate chamber. c-d
Umbilical views showing details of a peripheral spine. e Lateral view showing interiomarginal
aperture and keeled outline.
5. Adult specimen with sinistral coiling. a-c Spiral side, note the early nodose whorls and the
microperforate surface of the ultimate chamber. d Umbilical view showing strong nodose
ornamentation.
6. Adult dextral coiled specimen with peripheral spines. a Spiral view. b Umblical view.
c Lateral view.
Scale Bars: 100 µm
Plate B
Early ontogenetic stages of Pararotalia calcariformata
1.-3. Earliest stage specimens with three and four chambers. 1 Spiral view of specimen with
incipient peripheral pseudospines. 2 Spiral view of specimen with peripheral pseudospines. 3.
Umbilical view of specimens without peripheral pseudospines, showing incipient keel. Note the
pustulate surface.
4. Juvenile specimen with peripheral pseudospine on each chamber. a. Spiral view.
b Umbilical view, note the distinct trapezoidal shape of the chambers in the spiral and umbilical
sides. c Lateral view. d Details of the interiomarginal aperture.
5. Juvenile specimen with peripheral pseudospine on each chamber. a- b Spiral views showing
well-developed pore mounds and trapezoidal shaped chambers.
6. Juvenile specimen with peripheral pseudospine on each chamber. a- c Spiral views. d-f
Umbilical views, note the relatively large pore mounds on the surface both sides of the test. f
Details of the peripheral pseudospine.
Scale Bars: 1-3 = 10µm, 4-6d = 100µm and 6e = 50µm
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